Published September 26, 2025 | Version v1

Eutrophication boosts the competitive advantage of invasive gibel carp over endangered crucian carp

  • 1. Institute of Hydrobiology, Biology Centre of the Czech Academy of Sciences, České Budějovice, Czech Republic|University of South Bohemia, České Budějovice, Czech Republic
  • 2. Biology Centre, Czech Academy of Sciences, České Budějovice, Czech Republic|University of South Bohemia, České Budějovice, Czech Republic
  • 3. Czech University of Life Sciences Prague, Praha, Czech Republic
  • 4. Institute of Hydrobiology, Biology Centre of the Czech Academy of Sciences, České Budějovice, Czech Republic
  • 5. University of South Bohemia, České Budějovice, Czech Republic|Biology Centre, Czech Academy of Sciences, České Budějovice, Czech Republic|Institute of Hydrobiology, Biology Centre of the Czech Academy of Sciences, České Budějovice, Czech Republic
  • 6. University of South Bohemia, České Budějovice, Czech Republic|Biology Centre, Czech Academy of Sciences, České Budějovice, Czech Republic

Description

Anthropogenic eutrophication poses a significant threat to freshwater environments globally. It also influences the population dynamics of invasive and native species, yet the cumulative effects of eutrophication and invasive species on native organisms are not well understood. We used invasive gibel carp (Carassius gibelio Bloch, 1782) and native crucian carp (Carassius carassius Linnaeus, 1758) as model taxa to investigate how eutrophication influences their interspecific competition. Carassius carassius is being outcompeted by invasive C. gibelio across most of its native range, and eutrophication may play a role in the competitive displacement of C. carassius. We explored how varying eutrophication levels influence growth and survival of both species in laboratory and mesocosm experiments, where we exposed them to different feeding rations and nutrient levels, respectively. We hypothesized that (1) C. gibelio benefits more from increased nutrient levels than C. carassius—which favors invasive C. gibelio over native C. carassius in interspecific competition—and that (2) the growth and survival of C. carassius deteriorate under interspecific competition with C. gibelio compared to intraspecific competition. Our experiments revealed that excess nutrient availability was more beneficial to the invasive C. gibelio. Overall, C. gibelio individuals grew more than C. carassius, and the difference between the two species increased with nutrient levels in both experiments, especially in the weight of smaller individuals. Moreover, interspecific competition led to lower survival in C. carassius but not in C. gibelio in the mesocosm experiment. This study shows that eutrophication may modulate interspecific competition, and cultural eutrophication of freshwater habitats may enhance the success of invasive species.

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References

  • Adler PB, Smull D, Beard KH, Choi RT, Furniss T, Kulmatiski A, Meiners JM, Tredennick AT, Veblen KE (2018) Competition and coexistence in plant communities: Intraspecific competition is stronger than interspecific competition. Ecology Letters 21: 1319–1329. https://doi.org/10.1111/ele.13098
  • Alexander TJ, Vonlanthen P, Seehausen O (2017) Does eutrophication-driven evolution change aquatic ecosystems? Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 372: 20160041. https://doi.org/10.1098/rstb.2016.0041
  • Auwerx J, Wichelen J (2023) Unknown and still unloved: the story of the crucian carp (Carassius carassius) in Flanders. XVII European congress of ichthyology, Prague.
  • Aydin H, Gaygusuz Ö, Serhan TA, Top N, Emiroğlu Ö, Gürsoy GÇ (2011) Invasion of freshwater bodies in the Marmara region (northwestern Turkey) by nonnative gibel carp, Carassius gibelio (Bloch, 1782). Turkish Journal of Zoology 35: 829–836. https://doi.org/10.3906/zoo-1007-31
  • Blumenthal DM (2006) Interactions between resource availability and enemy release in plant invasion. Ecology Letters 9: 887–895. https://doi.org/10.1111/j.1461-0248.2006.00934.x
  • Bolker B (2023) bbmle: Tools for General Maximum Likelihood Estimation. https://cran.r-project.org/web/packages/bbmle
  • Bondarev D, Fedushko M, Hubanova N, Novitskiy R, Kunakh O, Zhukov O (2023) Temporal dynamics of the fish communities in the reservoir: The influence of eutrophication on ecological guilds structure. Ichthyological Research 70: 21–39. https://doi.org/10.1007/s10228-021-00854-x
  • Brönmark C, Miner JG (1992) Predator-induced phenotypical change in body morphology in crucian carp. Science 258: 1348–1350. https://doi.org/10.1126/science.258.5086.1348
  • Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg WC, Nielsen A, Skaug HJ, Machler M, Bolker BM (2017) glmmTMB balances speed and flexibility among packages for Zero-inflated Generalized Linear Mixed Modeling. The R Journal 9: 378–400. https://doi.org/10.32614/RJ-2017-066
  • Burnham KP, Anderson DR (2002) A practical information-theoretic approach. Model Selection and Multimodel Inference 2: 70–71.
  • Busst GMA, Britton JR (2017) Comparative trophic impacts of two globally invasive cyprinid fishes reveal species-specific invasion consequences for a threatened native fish. Freshwater Biology 62: 1587–1595. https://doi.org/10.1111/fwb.12970
  • Byers JE (2002) Impact of non‐indigenous species on natives enhanced by anthropogenic alteration of selection regimes. Oikos 97: 449–458. https://doi.org/10.1034/j.1600-0706.2002.970316.x
  • Carosi A, Lorenzoni F, Lorenzoni M (2023) Synergistic Effects of Climate Change and Alien Fish Invasions in Freshwater Ecosystems: A Review. Fishes 8: 486. https://doi.org/10.3390/fishes8100486
  • Copp GH, Bianco PG, Bogutskaya NG, Eros T, Falka I, Ferreira MT, Fox MG, Freyhof J, Gozlan RE, Grabowska J, Kováč V, Moreno-Amich R, Naseka AM, Peňáz M, Povž M, Przybylski M, Robillard M, Russell IC, Stakenas S, Šumer S, Vila-Gispert A, Wiesner C (2005) To be, or not to be, a non-native freshwater fish? Journal of Applied Ichthyology 21: 242–262. https://doi.org/10.1111/j.1439-0426.2005.00690.x
  • Crivelli AJ (1995) Are fish introductions a threat to endemic freshwater fishes in the northern Mediterranean region? Biological Conservation 72: 311–319. https://doi.org/10.1016/0006-3207(94)00092-5
  • David P, Thébault E, Anneville O, Duyck P-F, Chapuis E, Loeuille N (2017) Impacts of invasive species on food webs: A review of empirical data. Advances in Ecological Research 56: 1–60. https://doi.org/10.1016/bs.aecr.2016.10.001
  • Davis MA, Grime JP, Thompson K (2000) Fluctuating resources in plant communities: A general theory of invasibility. Journal of Ecology 88: 528–534. https://doi.org/10.1046/j.1365-2745.2000.00473.x
  • de Meo I, Østbye K, Kahilainen KK, Poléo ABS (2023) The role of predation risk in structuring life-history traits of crucian carp (Carassius carassius) in a series of small boreal lakes. Journal of Fish Biology 103(5): 939–949. https://doi.org/10.1111/jfb.15485
  • Dijoux S, Pichon NA, Sentis A, Boukal DS (2024) Body size and trophic position determine the outcomes of species invasions along temperature and productivity gradients. Ecology Letters 27: e14310. https://doi.org/10.1111/ele.14310
  • Directive H (1992) Council Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and flora. Official Journal of the European Union 206: 50.
  • Dmitriew CM (2011) The evolution of growth trajectories: What limits growth rate? Biological Reviews of the Cambridge Philosophical Society 86: 97–116. https://doi.org/10.1111/j.1469-185X.2010.00136.x
  • Dodds W, Smith V (2016) Nitrogen, phosphorus, and eutrophication in streams. Inland Waters 6: 155–164. https://doi.org/10.5268/IW-6.2.909
  • El-Sheekh M, Abdel-Daim MM, Okba M, Gharib S, Soliman A, El-Kassas H (2021) Green technology for bioremediation of the eutrophication phenomenon in aquatic ecosystems: A review. African Journal of Aquatic Science 46: 274–292. https://doi.org/10.2989/16085914.2020.1860892
  • Elgin EL, Tunna HR, Jackson LJ (2014) First confirmed records of Prussian carp, Carassius gibelio (Bloch, 1782) in open waters of North America. BioInvasions Records 3: 275–282. https://doi.org/10.3391/bir.2014.3.4.09
  • Emiroğlu Ö, Tarkan AS, Top N, Başkurt S, Sülün Ş (2012) Growth and Life History Traits of a Highly Exploited Population of Non-Native Gibel carp, Carassius gibelio from a Large Eutrophic Lake (Lake Uluabat, NW Turkey): Is Reproduction the Key Factor for Establishment Success? Turkish Journal of Fisheries and Aquatic Sciences 12(4). https://doi.org/10.4194/1303-2712-v12_4_20
  • Eurich JG, McCormick MI, Jones GP (2018) Direct and indirect effects of interspecific competition in a highly partitioned guild of reef fishes. Ecosphere 9: e02389. https://doi.org/10.1002/ecs2.2389
  • European Environment Agency (EEA) (2018) European waters - assessment of status and pressures 2018: 1–85. https://doi.org/10.2800/303664
  • Fernández-Alías A, Montaño-Barroso T, Conde-Caño M-R, Manchado-Pérez S, López-Galindo C, Quispe-Becerra J-I, Marcos C, Pérez-Ruzafa A (2022) Nutrient overload promotes the transition from top-down to bottom-up control and triggers dystrophic crises in a Mediterranean coastal lagoon. The Science of the Total Environment 846: 157388. https://doi.org/10.1016/j.scitotenv.2022.157388
  • Fuller RC (2016) Editorial Reconciling concepts, theory, and empirical patterns surrounding cascade reinforcement. Current Zoology 62: 131–134. https://doi.org/10.1093/cz/zow011
  • Gallardo B, Clavero M, Sánchez MI, Vilà M (2016) Global ecological impacts of invasive species in aquatic ecosystems. Global Change Biology 22: 151–163. https://doi.org/10.1111/gcb.13004
  • Gaygusuz Ö, Tarkan AS, Gaygusuz ÇG (2007) Changes in the fish community of the Ömerli Reservoir (Turkey) following the introduction of non-native gibel carp Carassius gibelio (Bloch, 1782) and other human impacts. Aquatic Invasions 2: 117–120. https://doi.org/10.3391/ai.2007.2.2.6
  • Gido KB, Franssen NR (2007) Invasion of stream fishes into low trophic positions. Ecology Freshwater Fish 16: 457–464. https://doi.org/10.1111/j.1600-0633.2007.00235.x
  • González AL, Kominoski JS, Danger M, Ishida S, Iwai N, Rubach A (2010) Can ecological stoichiometry help explain patterns of biological invasions? Oikos 119: 779–790. https://doi.org/10.1111/j.1600-0706.2009.18549.x
  • Gozlan RE, Britton JR, Cowx I, Copp GH (2010) Current knowledge on non-native freshwater fish introductions. Journal of Fish Biology 76: 751–786. https://doi.org/10.1111/j.1095-8649.2010.02566.x
  • Gu Q, Wang S, Zhong H, Yuan H, Yang J, Yang C, Huang X, Xu X, Wang Y, Wei Z, Wang J, Liu S (2022) Phylogeographic relationships and the evolutionary history of the Carassius auratus complex with a newly born homodiploid raw fish (2nNCRC). BMC Genomics 23: 242. https://doi.org/10.1186/s12864-022-08468-x
  • Gurevitch J, Morrow LL, Wallace A, Walsh JS (1992) A Meta-Analysis of Competition in Field Experiments. American Naturalist 140: 539–572. https://doi.org/10.1086/285428
  • Hartig F (2020) DHARMa: residual diagnostics for hierarchical (multi-level / mixed) regression models. R package version 0.3.
  • Havel JE, Kovalenko KE, Thomaz SM, Amalfitano S, Kats LB (2015) Aquatic invasive species: Challenges for the future. Hydrobiologia 750: 147–170. https://doi.org/10.1007/s10750-014-2166-0
  • Hensel K (1971) Some notes on the systematic status of Carassius auratus gibelio (Bloch, 1782) with further record of this fish from the Danube River in Czechoslovakia. Věstník Československé Společnosti Zoologické.
  • Holopainen IJ, Tonn WM, Paszkowski CA (1997) Tales of two fish: The dichotomous biology of crucian carp (Carassius carassius (L.)) in northern Europe. Annales Zoologici Fennici 34: 1–22.
  • Inouye BD (2001) Response surface experimental designs for investigating interspecific competition. Ecology 82: 2696–2706. https://doi.org/10.1890/0012-9658(2001)082[2696:RSEDFI]2.0.CO;2
  • Jeffries DL, Copp GH, Lawson Handley L, Olsén KH, Sayer CD, Hänfling B (2016) Comparing RADseq and microsatellites to infer complex phylogeographic patterns, an empirical perspective in the Crucian carp, Carassius carassius, L. Molecular Ecology 25: 2997–3018. https://doi.org/10.1111/mec.13613
  • Jobling M, Baardvik BM (1994) The influence of environmental manipulations on inter– and intra–individual variation in food acquisition and growth performance of Arctic charr, Salvelinus alpinus. Journal of Fish Biology 44. https://doi.org/10.1111/j.1095-8649.1994.tb01277.x
  • Kalous L, Rylková K, Bohlen J, Šanda R, Petrtýl M (2013) New mtDNA data reveal a wide distribution of the Japanese ginbuna Carassius langsdorfii in Europe. Journal of Fish Biology 82: 703–707. https://doi.org/10.1111/j.1095-8649.2012.03492.x
  • Kauppi L, Norkko J, Ikonen J, Norkko A (2017) Seasonal variability in ecosystem functions: Quantifying the contribution of invasive species to nutrient cycling in coastal ecosystems. Marine Ecology Progress Series 572: 193–207. https://doi.org/10.3354/meps12171
  • Khan MN, Mohammad F (2014) Eutrophication: Challenges and Solutions. In: Eutrophication: Causes, Consequences and Control. Springer Netherlands, Dordrecht, 1–15. https://doi.org/10.1007/978-94-007-7814-6_1
  • Kolar V, Boukal DS (2020) Habitat preferences of the endangered diving beetle Graphoderus bilineatus: Implications for conservation management. Insect Conservation and Diversity 13: 480–494. https://doi.org/10.1111/icad.12433
  • Kolar V, Tichanek F, Tropek R (2021) Evidence-based restoration of freshwater biodiversity after mining: Experience from Central European spoil heaps. Journal of Applied Ecology 58: 1921–1932. https://doi.org/10.1111/1365-2664.13956
  • Kolar V, Francová K, Vrba J, Grill S, Boukal DS (2023) Widespread long-term declines of littoral areas in protected and unprotected Czech fishponds. Ecological Engineering 194: 107042. https://doi.org/10.1016/j.ecoleng.2023.107042
  • Kopáček J, Hejzlar J (1993) Semi-Micro Determination of Total Phosphorus in Fresh Waters with Perchloric Acid Digestion. International Journal of Environmental Analytical Chemistry 53: 173–183. https://doi.org/10.1080/03067319308045987
  • Kottelat M, Freyhof J (2007) Handbook of European freshwater fishes (Vol. 13). Publications Kottelat, Cornol, Switzerland.
  • Lancaster HF, Drenner RW (1990) Experimental Mesocosm Study of the Separate and Interaction Effects of Phosphorus and Mosquitofish (Gambusia affinis) on Plankton Community Structure. Canadian Journal of Fisheries and Aquatic Sciences 47: 471–479. https://doi.org/10.1139/f90-051
  • Leger EA, Espeland EK (2010) Perspective: Coevolution between native and invasive plant competitors: implications for invasive species management. Evolutionary Applications 3: 169–178. https://doi.org/10.1111/j.1752-4571.2009.00105.x
  • Lekevičius E (2009) Vacant niches in nature, ecology, and evolutionary theory: A mini-review. Ekologija (Lietuvos Mokslu Akademija) 55: 165–174.
  • Lüdecke D (2018) ggeffects: Tidy Data Frames of Marginal Effects from Regression Models. Journal of Open Source Software 3: 772. https://doi.org/10.21105/joss.00772
  • Lüdecke D, Ben-Shachar M, Patil I, Makowski D (2020) Extracting, Computing and Exploring the Parameters of Statistical Models using R. Journal of Open Source Software 5: 2445. https://doi.org/10.21105/joss.02445
  • Meller SA, Crowl TA (2006) Effects of common carp (Cyprinus carpio) on macrophytes and invertebrate communities in a shallow lake. Freshwater Biology 51: 85–94. https://doi.org/10.1111/j.1365-2427.2005.01477.x
  • Milardi M, Aschonitis V, Gavioli A, Lanzoni M, Fano EA, Castaldelli G (2018) Run to the hills: Exotic fish invasions and water quality degradation drive native fish to higher altitudes. The Science of the Total Environment 624: 1325–1335. https://doi.org/10.1016/j.scitotenv.2017.12.237
  • Milardi M, Iemma A, Waite IR, Gavioli A, Soana E, Castaldelli G (2022) Natural and anthropogenic factors drive large-scale freshwater fish invasions. Scientific Reports 12: 1–12. https://doi.org/10.1038/s41598-022-14556-5
  • Mooney HA, Cleland EE (2001) The evolutionary impact of invasive species. Proceedings of the National Academy of Sciences of the United States of America 98: 5446–5451. https://doi.org/10.1073/pnas.091093398
  • Navodaru I, Buijse AD, Staras M (2002) Effects of Hydrology and Water Quality on the Fish Communityin Danube Delta Lakes. International Review of Hydrobiology 87: 329–348. https://doi.org/10.1002/1522-2632(200205)87:2/3<329:AID-IROH329>3.0.CO;2-J
  • Ohlberger J (2013) Climate warming and ectotherm body size – from individual physiology to community ecology. Functional Ecology 27: 991–1001. https://doi.org/10.1111/1365-2435.12098
  • Özdilek SY, Jones RI (2014) The Diet Composition and Trophic Position of Introduced Prussian Carp Carassius gibelio (Bloch, 1782) and Native Fish Species in a Turkish River. Turkish Journal of Fisheries and Aquatic Sciences 14: 769–776. https://doi.org/10.4194/1303-2712-v14_3_19
  • Papoušek I, Vetešník L, Halačka K, Lusková V, Humpl M, Mendel J (2008) Identification of natural hybrids of gibel carp Carassius auratus gibelio (Bloch) and crucian carp Carassius carassius (L.) from lower Dyje River floodplain (Czech Republic). Journal of Fish Biology 72: 1230–1235. https://doi.org/10.1111/j.1095-8649.2007.01783.x
  • Penning WE, Mjelde M, Dudley B, Hellsten S, Hanganu J, Kolada A, van den Berg M, Poikane S, Phillips G, Willby N, Ecke F (2008) Classifying aquatic macrophytes as indicators of eutrophication in European lakes. Aquatic Ecology 42: 237–251. https://doi.org/10.1007/s10452-008-9182-y
  • Pentyliuk N, Schmidt B, Poesch MS, Green SJ (2023) Recreational angler reporting as a tool for tracking the distribution of invasive Prussian carp (Carassius gibelio). Conservation Science and Practice 5: e12850. https://doi.org/10.1111/csp2.12850
  • Prepas EE, Charette T (2003) Worldwide Eutrophication of Water Bodies: Causes, Concerns, Controls. Treatise on Geochemistry 311–331. https://doi.org/10.1016/B0-08-043751-6/09169-6
  • Preston DL, Hedman HD, Johnson PTJ (2018) Nutrient availability and invasive fish jointly drive community dynamics in an experimental aquatic system. Ecosphere 9: e02153. https://doi.org/10.1002/ecs2.2153
  • R Core Team (2024) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/
  • Razlutskij V, Mei X, Maisak N, Sysova E, Lukashanets D, Makaranka A, Jeppesen E, Zhang X (2021) Omnivorous Carp (Carassius gibelio) Increase Eutrophication in Part by Preventing Development of Large-Bodied Zooplankton and Submerged Macrophytes. Water 13: 1497. https://doi.org/10.3390/w13111497
  • Ribeiro F, Rylková K, Moreno-Valcárcel R, Carrapato C, Kalous L (2015) Prussian carp Carassius gibelio: A silent invader arriving to the Iberian Peninsula. Aquatic Ecology 49: 99–104. https://doi.org/10.1007/s10452-015-9508-5
  • Richardson MJ, Whoriskey FG, Roy LH (1995) Turbidity generation and biological impacts of an exotic fish Carassius auratus, introduced into shallow seasonally anoxic ponds. Journal of Fish Biology 47: 576–585. https://doi.org/10.1111/j.1095-8649.1995.tb01924.x
  • Rolton A, Rhodes L, Hutson KS, Biessy L, Bui T, MacKenzie L, Symonds JE, Smith KF (2022) Effects of Harmful Algal Blooms on Fish and Shellfish Species: A Case Study of New Zealand in a Changing Environment. Toxins 14: 341. https://doi.org/10.3390/toxins14050341
  • Rosset V, Angélibert S, Arthaud F, Bornette G, Robin J, Wezel A, Vallod D, Oertli B (2014) Is eutrophication really a major impairment for small waterbody biodiversity? Journal of Applied Ecology 51: 415–425. https://doi.org/10.1111/1365-2664.12201
  • Rylková K, Kalous L, Bohlen J, Lamatsch DK, Petrtýl M (2013) Phylogeny and biogeographic history of the cyprinid fish genus Carassius (Teleostei: Cyprinidae) with focus on natural and anthropogenic arrivals in Europe. Aquaculture (Amsterdam, Netherlands) 380–383: 13–20. https://doi.org/10.1016/j.aquaculture.2012.11.027
  • Sayer CD, Copp GH, Emson D, Godard MJ, Zięba G, Wesley KJ (2011) Towards the conservation of crucian carp Carassius carassius: Understanding the extent and causes of decline within part of its native English range. Journal of Fish Biology 79: 1608–1624. https://doi.org/10.1111/j.1095-8649.2011.03059.x
  • Sayer CD, Emson D, Patmore IR, Greaves HM, West WP, Payne J, Davies GD, Tarkan AS, Wiseman G, Cooper B, Grapes T, Cooper G, Copp GH (2020) Recovery of the crucian carp Carassius carassius (L.): Approach and early results of an English conservation project. Aquatic Conservation 30: 2240–2253. https://doi.org/10.1002/aqc.3422
  • Schindler DW (2006) Recent advances in the understanding and management of eutrophication. Limnology and Oceanography 51: 356–363. https://doi.org/10.4319/lo.2006.51.1_part_2.0356
  • Seehausen O, Van Alphen JJM, Witte F (1997) Cichlid fish diversity threatened by eutrophication that curbs sexual selection. Science 277: 1808–1811. https://doi.org/10.1126/science.277.5333.1808
  • Simberloff D (2005) Non-native Species DO Threaten the Natural Environment! Journal of Agricultural & Environmental Ethics 18: 595–607. https://doi.org/10.1007/s10806-005-2851-0
  • Šmejkal M, Bartoň D, Duras J, Horký P, Muška M, Kubečka J, Pfauserová N, Tesfaye M, Slavík O (2023) Living on the edge: Reservoirs facilitate enhanced interactions among generalist and rheophilic fish species in tributaries. Frontiers in Environmental Science 11: 1099030. https://doi.org/10.3389/fenvs.2023.1099030
  • Šmejkal M, Thomas K, Kořen V, Kubečka J (2024) The 50-year history of anglers' record catches of genus Carassius: Circumstantial evidence of wiping out the native species by invasive conspecific. NeoBiota 92: 111–128. https://doi.org/10.3897/neobiota.92.121288
  • Šmejkal M, Kalous L, Auwerx J, Gorule PA, Jarić I, Dočkal O, Fedorčák J, Muška M, Thomas K, Takács P, Ferincz Á, Choleva L, Lamatsch DK, Wanzenböck J, Van Wichelen J (2025) Wetland fish in peril: A synergy between habitat loss and biological invasions drives the extinction of neglected native fauna. Biological Conservation 302: 110948. https://doi.org/10.1016/j.biocon.2024.110948
  • Smith VH (2003) Eutrophication of freshwater and coastal marine ecosystems a global problem. Environmental Science and Pollution Research International 10: 126–139. https://doi.org/10.1065/espr2002.12.142
  • Smith VH, Schindler DW (2009) Eutrophication science: Where do we go from here? Trends in Ecology & Evolution 24: 201–207. https://doi.org/10.1016/j.tree.2008.11.009
  • Smith VH, Tilman GD, Nekola JC (1999) Eutrophication: Impacts of excess nutrient inputs on freshwater, marine, and terrestrial ecosystems. Environmental Pollution 100: 179–196. https://doi.org/10.1016/S0269-7491(99)00091-3
  • Souza AT, Argillier C, Blabolil P, Děd V, Jarić I, Monteoliva AP, Reynaud N, Ribeiro F, Ritterbusch D, Sala P, Šmejkal M, Volta P, Kubečka J (2022) Empirical evidence on the effects of climate on the viability of common carp (Cyprinus carpio) populations in European lakes. Biological Invasions 24: 1213–1227. https://doi.org/10.1007/S10530-021-02710-5
  • Stendera S, Adrian R, Bonada N, Cañedo-Argüelles M, Hugueny B, Januschke K, Pletterbauer F, Hering D (2012) Drivers and stressors of freshwater biodiversity patterns across different ecosystems and scales: A review. Hydrobiologia 696: 1–28. https://doi.org/10.1007/s10750-012-1183-0
  • Sterner RW, Elser JJ (2003) Ecological stoichiometry: the biology of elements from molecules to the biosphere. Princeton University Press. https://doi.org/10.1515/9781400885695
  • Szczerbowski JA (2002) Carassius auratus (Linnaeus, 1758). In: Banarescu P, Paepke HJ (Eds) The freshwater fishes of Europe, Vol. 5/III, Cyprinidae 2. AULA-Verlag, Wiesbaden, 5–41.
  • Tapkir S, Boukal D, Kalous L, Bartoň D, Souza AT, Kolar V, Soukalová K, Duchet C, Gottwald M, Šmejkal M (2022) Invasive gibel carp (Carassius gibelio) outperforms threatened native crucian carp (Carassius carassius) in growth rate and effectiveness of resource use: Field and experimental evidence. Aquatic Conservation 32: 1901–1912. https://doi.org/10.1002/aqc.3894
  • Tapkir S, Thomas K, Kalous L, Vašek M, Meador TB, Šmejkal M (2023) Invasive gibel carp use vacant space and occupy lower trophic niche compared to endangered native crucian carp. Biological Invasions 29: 2917–2928. https://doi.org/10.1007/s10530-023-03081-9
  • Tarkan AS, Gaygusuz O, Gürsoy Gaygusuz C, Saç G, Copp GH (2012a) Circumstantial evidence of gibel carp, Carassius gibelio, reproductive competition exerted on native fish species in a mesotrophic reservoir. Fisheries Management and Ecology 19: 167–177. https://doi.org/10.1111/j.1365-2400.2011.00839.x
  • Tarkan AS, Copp GH, Top N, Özdemir N, Önsoy B, Bilge G, Filiz H, Yapici S, Ekmekçi FG, Kirankaya ŞG, Emiroǧlu O, Gaygusuz O, C. GG, Oymak A, Özcan G, Saç G (2012b) Are introduced gibel carp Carassius gibelio in Turkey more invasive in artificial than in natural waters? Fisheries Management and Ecology 19: 178–187. https://doi.org/10.1111/j.1365-2400.2011.00841.x
  • Tarkan AS, Almeida D, Godard MJ, Gaygusuz Ö, Rylands M, Sayer CD, Zieba G, Copp GH (2016) A review and meta-analysis of growth and life-history traits of a declining European freshwater fish, crucian carp Carassius carassius. Aquatic Conservation 26: 212–224. https://doi.org/10.1002/aqc.2580
  • Taylor JM, King RS, Pease AA, Winemiller KO (2014) Nonlinear response of stream ecosystem structure to low‐level phosphorus enrichment. Freshwater Biology 59: 969–984. https://doi.org/10.1111/fwb.12320
  • ter Braak CJF, Smilauer P (2012) Canoco reference manual and user's guide: software for ordination, version 5.0. Microcomputer Power, Ithaca, USA, 343–354. https://doi.org/10.2/JQUERY.MIN.JS
  • Tesfaye M, Kumkar P, Hailu M, Verma CR, Tesfaye G, Petrtýl M, Getahun A, Šmejkal M, Kalous L (2025) Discovery of Japanese gengorobuna Carassius cuvieri Temminck & Schlegel, 1846 in Ethiopia: Implications for fisheries governance. BioInvasions Records 14: 155–167. https://doi.org/10.3391/BIR.2025.14.1.13
  • Thomas K, Brabec M, Kalous L, Gottwald M, Bartoň D, Grill S, Kořen V, Tapkir S, Šmejkal M (2024) Anthropogenic induced drivers of fish assemblages in small water bodies and conservation implications. Ecohydrology & Hydrobiology 25(3): 691–699. https://doi.org/10.1016/j.ecohyd.2024.11.003
  • Tibbets TM, Krist AC, Hall RO, Riley LA (2010) Phosphorus-mediated changes in life history traits of the invasive New Zealand mudsnail (Potamopyrgus antipodarum). Oecologia 163: 549–559. https://doi.org/10.1007/s00442-009-1522-7
  • Tilman D (1982) Resource competition and community structure. Princeton University Press, Princeton, NJ. https://doi.org/10.1515/9780691209654
  • Tsoumani M, Liasko R, Moutsaki P, Kagalou I, Leonardos I (2006) Length-weight relationships of an invasive cyprinid fish (Carassius gibelio) from 12 Greek lakes in relation to their trophic states. Journal of Applied Ichthyology 22: 281–284. https://doi.org/10.1111/j.1439-0426.2006.00768.x
  • Vadeboncoeur Y, Moore MV, Stewart SD, Chandra S, Atkins KS, Baron JS, Bouma-Gregson K, Brothers S, Francoeur SN, Genzoli L, Higgins SN, Hilt S, Katona LR, Kelly D, Oleksy IA, Ozersky T, Power ME, Roberts D, Smits AP, Timoshkin O, Tromboni F, Vander Zanden MJ, Volkova EA, Waters S, Wood SA, Yamamuro M (2021) Blue Waters, Green Bottoms: Benthic Filamentous Algal Blooms Are an Emerging Threat to Clear Lakes Worldwide. Bioscience 71: 1011–1027. https://doi.org/10.1093/biosci/biab049
  • Van den Brink PJ, Ter Braak CJF (1999) Principal response curves: Analysis of time‐dependent multivariate responses of biological community to stress. Environmental Toxicology and Chemistry 18: 138–148. https://doi.org/10.1002/etc.5620180207
  • Vašek M, Prchalová M, Říha M, Blabolil P, Čech M, Draštík V, Frouzová J, Jůza T, Kratochvíl M, Muška M, Peterka J, Sajdlová Z, Šmejkal M, Tušer M, Vejřík L, Znachor P, Mrkvička T, Seɱa J, Kubečka J (2016) Fish community response to the longitudinal environmental gradient in Czech deep-valley reservoirs: Implications for ecological monitoring and management. Ecological Indicators 63: 219–230. https://doi.org/10.1016/j.ecolind.2015.11.061
  • Vilizzi L, Tarkan AS, Copp GH (2015) Experimental Evidence from Causal Criteria Analysis for the Effects of Common Carp Cyprinus carpio on Freshwater Ecosystems: A Global Perspective. Reviews in Fisheries Science & Aquaculture 23: 253–290. https://doi.org/10.1080/23308249.2015.1051214
  • Wang X, Cui Z, Guo Q, Han X, Wang J (2009) Distribution of nutrients and eutrophication assessment in the Bohai Sea of China. Chinese Journal of Oceanology and Limnology 27: 177–183. https://doi.org/10.1007/s00343-009-0177-x
  • Wouters J, Janson S, Lusková V, Olsén KH (2012) Molecular identification of hybrids of the invasive gibel carp Carassius auratus gibelio and crucian carp Carassius carassius in Swedish waters. Journal of Fish Biology 80: 2595–2604. https://doi.org/10.1111/j.1095-8649.2012.03312.x
  • Zuur, (2009) Mixed Effects Models and Extensions in Ecology with R. Springer Verlag. https://doi.org/10.1007/978-0-387-87458-6